Professional Documents
Culture Documents
COMPARATIVE PHYSIOLOGY
TEXT-BOOKS OF ANIMAL
BIOLOGY
Edited by Professor Julian S. Huxley
Other volumes in preparation.
31
mi
COMPARATIVE
PHYSIOLOGY
BY
LANCELOT T. HOGBEN
M.A.(Cantab.), D.Sc.(Lond.)
Assistant Professor in Zoology, McGili» UNiVERiiTY
LONDON
SIDGWICK & JACKSON, LTD.
1926
PRINTED IN GREAT BRITAIN BY
WILLIAM CLOWES AND SONS, LIMITED
LONDON AND BECCLES.
TO
E. A. S. S.
AUTHOR'S PREFACE
There is, so far as I know, no work in English which aims at
medical studies.
My thanks are due to Dr. A. D. Macdonald and Mr. A. D.
Hobson, who read the MS., and to Professor Julian Huxley foe
valuable suggestions.
LANCELOT T. HOGBEN.
March i, 1925.
CONTENTS
CHAPTERS I— III
PAGE
Response —the
I. .......
Muscular Contraction
Manifestations of Vital Activity
i
II.
CHAPTERS IV—VI
Metabolism— THE Sources of Vital Energy
IV.
V.
Respiration
Nutrition
......... 64
85
VI. The Circulation of Body Fluids . . . . , loi
CHAPTERS VII- IX
Co-ordination — the Integration of Vital Activities
VII. Endocrine Co-ordination 118
VIII. The Mechanism of Nervous Conduction and Excitation . 134
XI. The Analysis of Behaviour in Animals . . . • 151
CHAPTERS X— XII
Reproduction — the Building up of a New
Animate Unit
X. The ......
.........
Fertilisation of the Egg 169
XI.
XII.
Inheritance
The Physiology .....
of Development
182
200
LIST OF ILLUSTRATIONS
1.
2.
Diagram
contraction
Scheme to illustrate
........
illustrating liberation of potential
.
6
8.
Effect of removal
Diagram
of calcium
24
10.
ciha to temperature ......
Relation of oxygen consumption and mechanical activity of
29
amoeboid movement
32
38
78
21. Relation of
CO2 pressure .......
amount of COj taken up by mammahan blood to
80
XIV LIST OF ILLUSTRATIONS
28.
brain .....
27. Inhibition of heart-beat of Limulus by electric stimulation of
Heart of Cephalopod
37. Excitability by
claw of Astacus ....
single or double stimuli of abductor nerve of
MUSCULAR CONTRACTION
and it is only within recent years that the work of A. V. Hill has
placed the question on a satisfactory basis.
{a) Mechanical Response in Muscle.— In attempting to
grasp the significance of the chemical and thermal aspects of
contraction, it is necessary to measure the potential mechanical
thus does not follow that the work it may be made to perform
it
In just the same way, the energy which we get out of a weighted
muscle is not simply the amount of energy liberated by the
contractile mechanism sensu stricto.
The way in which we may calculate the maximal amount of
work which the muscle capable of doing in virtue of the
is
W= r T.dl
(E being the extended length and C contracted length of the
muscle.)
In this form it is not possible to evaluate W directly, since
Fig. I.
it exerts no tension on the lever, i.e. till the clamp has been
the muscle can contract and relax in the total absence of oxygen,
though the presence of oxygen delays the onset of fatigue ;
Heat production in Ng
(make) ;the myograph the string of the galvanometer signal for the
; ;
myograph vertically being equal to 500 gms. tension. String tension 5 mm.
per m. V. the magnification being 285.
, Stimuli delivered to the cut nerve,
the cathode being at a point I'g cm. from the entry of the nerve into the
muscle; stimuli just-maximal induction shocks. Initial tension 90 gms.
Frequency of the myograph 460 per sec.
chemistry.
There is Httle doubt that the demarcation current arises
from the distribution of electrolytes in the muscle system its ;
Hke a vice. But if the foreign body is then made to slide out
of position, the valves remain set fast at the same degree of
closure. Uexkull (19 12) has shown that on cutting away the
smooth muscle, the remainder can be excited to contraction
by nervous stimulation but the contraction of the striated
;
1 COMPARATIVE PHYSIOLOGY
by its sustained tonus. A double neuromuscular mechanism
probably of the same type exists in the adductor of Astacus,
Homarus, Carcinus and other decapod Crustacea (Lapicque,
Keith Lucas). The economy of such an arrangement, which
combines the rapidity of action of striped muscle with the low
energy output of the tonus mechanism, is evident. Tonus is
not associated with increased metabolism, and cannot there-
fore be of the same nature as a low-grade tetanus.
The tonus of a catch muscle may be looked upon as a natural
form of isometric response. We are probably dealii(ig here not
with a change in the intensity factor but with the capacity factor
of the surface energy of the cell. It is possible to think of a
mechanism of isometric response by which a mechanical stress
sets up in some part of the protoplasmic system a change in
phase relations of the colloidal constituents such as to oppose it
by a virtual tension co- extensive with the maintenance of the
external force. The coagulation of liquid silk in stretching is
possibly analogous.
(g) Relation of the Muscle Cell to Electrolytes.— Certain
phenomena, notably those associated with the maintenance of
tone which has just been discussed, raise difficulty in the way
of any attempt to extend to plain muscle the conclusions
respecting the contractile mechanism in striped muscle. This
is also the case in considering the role of the hydrogen and
the other hand, found both for the cardiac muscle of the
it is
other.
On the whole the indications of recent work are distinctly
favourable to the role which Mines postulates for calcium.
The effects of the other polyvalent ions are more obscure, since
Mg . . , Ce . . . , La . . . , etc., agree with Ca in depressing the
striped muscle of the vertebrate, and that of the crustacean
heart ; while their action is opposite to that of Ca in a large
number of instances in the case of cardiac and plain muscle.
Thus in Pecten, removal of Mg, which is apparently essential
to the heart-beat, produces systolic arrest, while removal of
MUSCULAR CONTRACTION 19
^ __^mi*"'
Further Reading
24 COMPARATIVE PHYSIOLOGY
or recovery stroke. The form of the beat suggests that it is
during the rapid effective stroke that the cilium performs v^ork
on the surrounding medium. At the conclusion of the forward
movement, it can be seen that the ciHum, which at the beginning
of the effective stroke is a more or less rigid rod moving forward
on a pivot at its base, becomes limp a stress is set up w^hich
;
starts at its base and is transmitted thence to its free end. For
the purpose of forming a working hypothesis, it may be assumed,
as suggested by Gray, ''
that the energy which is expended by
the cilium is stored as tension energy." We are entitled to
surmise that this energy has
Forward or Effective Beat
its origin in some chemical
compound either in the cilium
itself or in the cell to which
it is The problem
attached.
to be faced is the elucidation
Backward or Recovery Beat of the sequence of events by
which chemical energy is con-
verted into kinetic energy
or as— implied above the —
chemical processes by which
Fig. 8. — Diagram of ciliary motion
(after Gray). the state of tension in the
cilium is relieved.
We shall here accept Gray's hypothesis as a basis for
discussion. The results of the foregoing survey of muscular
contraction have led us to conclude that the physical changes
which result in contraction are associated with the production
of lactic acidfrom carbohydrate without intake of oxygen, and
that oxygen is employed in the recovery process to restore the
mechanism to its original condition. Though ciliated
epithelium is structurally very different from muscle, there are
two sets of considerations which suggest the possibility that a
similar sequence of chemical phenomena might be found to
underlie the changes of physical state, which in both cases
result in liberating contractile energy. One is that lactic
acid an obligatory intermediary in the breakdown of carbo-
is
that the effect is a surface one, concerned only with the rate at
which the excitation stateis generated.
Acid.
CILIARY ACTIVITY 27
lacking.
Turning now to the second aspect of the problem, that is,
TEMPERATURE
Fig. 10. —The relation of oxygen consumption and mechanical activity of
cilia to temperature (after Gray).
HCl ....+
Citric O and Q
Tartaric ... ^
pH&O
Four velocity
curves obtained suc-
cessively from tlie
same Type B amoeba,.
Each curve taken
with a fresh acid, after
recovery. in "outside
sea water,"
30 ©c /
VELOCITVatr !
VELOCITYatIO' / .
30°C.
in solutions of
M
— concentration.
. .
repair does not occur ; the surface sets into a solid mass.
The antagonism between the monovalent and divalent ions is
M
in a solution of NaCl —
M
CaCl2 —
seen again in this process : .
or of KCl
13
. CaClo
^ —
208
b
416
z repair of a torn surface takes
^
place in the normal manner.
Pantin (1925) has also investigated the relation of electrolytes
to the velocity of amoeboid movement, and has obtained
somewhat analogous results. The marine amoeba is destroyed
in isotonic solutions of the chlorides of the four principal
kations present in sea water sodium, magnesium, potassium,
{i.e.
CILIARY ACTIVITY 37
a latent period of
about 5 sees, before contraction begins. Single make shocks
successively applied produce a summated effect if the inter-
is to say, dark brown prawns become deep blue and light ones
become pale blue. The nocturnal tint ordinarily persists till
CILIARY ACTIVITY 41
brown or red colour forms display " extension " of the red
pigment into the cell branches we should be tempted to
;
Further Reading
Chambers and Reznikoff. Proc. Soc. Exp. Biol. Med. 22, 1925.
Spaeth (19 16). Responses of Single Melanophores to Electrical Stimula-
tion. Am. Journ. Physiol. 41.
Keeble and Gamble (1904-6). The Colour Physiology of the Higher
Crustacea. Phil. Trans. Roy. Soc.
CHAPTER III
SECRETION
(2) in driving the filtrate along the narrow lumen of the tubules
at the observed rate of flow. From the first consideration it
follows that no secretion of urine can take place when the blood-
pressure falls to a value below the osmotic pressure of the
Total (non-protein)
Animal. |
Mammal (Homo)
Amphibian (Rana)
Teleost (Lophius)
Elasmobranch (Mustelus)
52 COMPARATIVE PHYSIOLOGY
tissues. According to Baglioni (1903) and Mines (1912),
addition of urea to the saline medium is essential to the mainte-
chemical entities.
The term " enzyme " was introduced by Kiihne (1878) to
obviate the confusion resulting from the use of the older term
" ferment," a term originally used to include the activities of
Frog serum injected into the body cavity of the crab specifi-
poisonous secretion of the
cally protects the latter against the
pedicellari^e of Echinoderms.
certain The mechanism of
immunity is exceedingly complex in addition to immunity
;
Further Reading
Swale Vincent. An Introduction to the Study of Secretion. Arnold.
Bernstein. Elektro biologic. Vieweg.
CusHNY. The Secretion of Urine. Longmans, Green.
Newton Har\^y. The Nature of Animal I>ight. Lippincott.
Calmette. Venoms. Bale & Danielsson.
CHAPTER IV
RESPIRATION
the lungs and gills with but little diminution of total respira-
tory activity. The following data from Krogh's (1904)
experiments show that the skin is a very important factor
in the respiration of the frog, especially when it is pointed
out that the total surface of the skin is only about one and
a half times the internal surface of the lungs.
66 COMPARATIVE PHYSIOLOGY
making use of the Bodo migrates to a
fact that the flagellate
region having a certain optimum oxygen Fox found tension.
that when pupae of Simulium, which respire by means of
filamentous appendages at the junction of head and thorax,
are placed in a suspension of Bodo, the micro-organisms collect
at first round the filaments in a dense congregation, and then
migrate outwards in a crescentic configuration, as the oxygen
concentration falls through absorption to a lower level than the
optimum. When Chironomus larv83 are similarly placed in
RESPIRATION 67
50
30
20
0;:GOj.
GOjv.
5^..
•• 10 15 20 25 30
5iO
I
I
5(8
I
57B
I
576 SifO
II and III give the spectrophotometric axes of the bands of the haemoglobin
of the horse (II) and Arenicola (III) (after Fox).
—
—\
(,10 5S0
IV
Am.S : V
mammalian haemoglobin (after Fox).
'
'
1 1
T70
RESPIRATION 8i
50 60 70
Fig. 22.
body fluid of Aplysia (Bottazzi) contains less than o"oi per cent,
of protein nitrogen. The isoelectric point of haemocyanin is
Further Reading
Books,
Barcroft's Respiratory Function of the Blood.
Krogh's Respiratory Exchange of Animals and Man.
Haldane's Organism and Environment.
Haldane's Respiration.
On Respiration in Cephalopods.
Polimanti (19 1 2). Beitrage zur physiologic von Sepia II. Arch. f. anat.
u. Physiol, p. 53 (1909)-
WiNTERSTEiN. Zur kenntnis der Blutgase wirbellose Seetiere. Biochem.
Zeitschr. 19.
Annelids.
Insects.
Fishes.
Pigments.
Carbon Dioxide.
Parsons and Parsons (1923). Transport of Carbon Dioxide in the Blood
of Some Marine Invertebrates. Journ. Gen. Physiol. 6.
CHAPTER V
NUTRITION
however, grew steadily, and the possibility that the fly larva
actually ingests the yeasts alone remained. On cultures of
compressed yeast-agar with yeast nucleo-proteins as the sole
88 COMPARATIVE PHYSIOLOGY
nitrogen supply, the sterile larvae were able to grow and pupate
normally. It thus appears that yeasts are the nitrogenous
food of Drosophila ; the simplest nutrient solution suitable
for the yeasts (and certain other micro-organisms) will replace
fermenting fruit in the ecology of Drosophila larvae. These
experiments of Baumberger are extremely suggestive in relation
to the diet of wood-boring animals, the significance of fungus
gardens, the curious habitat of such organisms as the vinegar
worm, and a host of other bionomic problems.
Researches have also been carried out on Drosophila
larvae in relation to the accessory food-factors or vitamins.
The term " vitamin " is one which at present can hardly be said
to convey more than a recognition of our failure to induce
mammals to grow healthily on a diet of purified carbohydrate,
fat, protein, etc., and our almost complete ignorance of those
NUTRITION 89
Digestive gland
stomach
style
heart
Fig. 24. —The crystalline style of the bivalve mollusc.
References
Bacot and Harden (1922). The Vitamin Requirements of Drosophila.
Biochem. Journ. 16.
plex one, taking place in some species in two stages. The first
stage in the clotting of crustacean blood corresponds more or
less to the coagulation process in that of the Arachnid, Limulus,
whose blood also clots rapidly. In Limulus the protein con-
tent of the blood is almost exclusively made up by the haemo-
cyanin and white blood corpuscles. Coagulation is essentially
a phenomenon of cytolysis (Alsberg and Clark), and can be
prevented by reagents which hinder cell agglutination. For
the first stage of coagulation in crustacean blood, which is
brought about by the cytolysis of special '' explosive " cells,
firstrecognised by Hardy (1892), immediately they come into
contact with foreign substance, it has been shown by L. Loeb
that calcium ions are not necessary, though they are necessary
for the second, which takes place in lobster blood about a
io6 COMPARATIVE PHYSIOLOGY
quarter of an hour later. This consists of a jellying of the
plasma, but according to Tait (191 8) it is accompanied by
a further cytolysis of corpuscles, which he designates thigmo-
cytes. Loeb states that lobster tissues yield specific coaguHns
for the blood of the same species. According to Tait all the
corpuscles of crustacean blood are actively phagocytic. The
subject has attracted a large number of investigators since
attention was focussed upon it by the pioneer labours of
Fredericq but much still remains to be done, especially
;
and Satacke (19 12) measured the pressure of the blood in the
abdominal aorta of the lobster by means of a water mano-
meter, using hirudin as anticoagulant, and found that with
the heart beating at a rate of 51 per minute the average
pressure-reading was only about 8 mm. of mercury.
The most extensive studies on the circulatory system of
any invertebrate are the investigations of Carlson on the heart
of the king-crab, Limulus. In this arachnid' the heart retains
12 11
10 9 S"7
7-8 Cardiac nerve from brain ; 9-1 1 cardiac nerve from abdominal
ganglia cardiac ganglion ;
p.n.c. lateral nerves.
iliiii__MAiJLLi^
Fig. 27.- -Inhibition of heart beat of Limulus by electrical stimulation
of brain with weak current (Carlson).
Ql0ll72I° 1*92
Qioi3723° 1*90
Q1015725'' 2' 20
Qioi9729° 2- 18
—
The Circulatory System in Molluscs. Among the Cepha-
lopod molluscs the circulatory system reaches the highest
degree of specialisation met with among the invertebrate
phyla. The anatomical relations are briefly as follows.
Venous blood is collected by the caval vein which bifurcates
to form two branches supplying the gills. These afferent
branchial vessels dilate at the base of the gills into rhythmically
contractile branchial hearts, which drive the colourless venous
blood through the gill-capillaries. Oxygenated blue blood is
collected by the efferent vessels directly to the auricles of the
systemic heart, the ventricle (which may be divided) pumping
the arterial blood into the aortae. This would seem to be a
much more efficient device for supplying oxygenated blood to
the tissues at high pressure than the arrangement which exists
in fishes, where the force of the heart's heat has to overcome
the resistance of the gill-capillaries as well as that of the body
THE CIRCULATION OF BODY FLUIDS in
capillaries.And it is an experimental fact that the blood-
pressure in the arterial system of the Cephalopod is much
higher than in that of the fish. The aortic pressure
of the octopus has been determined by several observers,
first by Fuchs, who found that it varied between 25-80 mm.
Hg ;
probably the more representative. But
latter figure is
the efficiency of the mechanism becomes even more striking
when the relation between the rhythm of the branchial hearts,
of the systemic hearts, and of the respiratory movements is
studied. The pulsation of the branchial hearts precedes
that of the systemic hearts, but at the same rate (about 30 beats
L. Aurfcle
Ventrfcle
into ill- defined lacunae which finally converge upon the gill-
sinuses. The total resistance of the peripheral circulation
is so small that the blood-pressure is not of the same order
I
:: — ...
.. ... . . . .
Cardium . . . 15-17 . .
,,
Gasteropods
Helix 53 Lang
Cephalopods
Octopus . . . . 55 . . . . . Bauer.
Sepia . . • • 35 • • • • • • Fry.
the base of the pharyngeal sac, so that one end lies dorsal to
the gut and may extend to the nerve gangUon, the other end
being ventral on the opposite side of the pharnyx. The heart-
rhythm is made up of alternating series of beats progressing
respectively from the dorsal (advisceral) and ventral (abvis-
ceral) ends of the heart. In Ascidia mentula, as described by
Day (1921), each pulsation-series lasts from two to four
THE CIRCULATION OF BODY FLUIDS 115
Abvisceral 17 15 36 36
Advisceral 44 51 65
References
Alsberg (19 1 4). The Proteins of the Blood of Limulus. J. Biol. Chem.
19.
Brucke and Satacke (1912). Der Arterielle Blutdruck des Hummers.
Zeitschr. Allg. Physiol. 7.
Dakin (1908). The Osmotic Concentration of the Blood of Fishes, etc.
Biochem. J. 3.
Carlson (1904-5). Nervous Origin of the Heart Beat in Limulus. Am.
Journ. Physiol. 12.
(1906). Chemical Conditions of Heart Action in Limulus. Ibid. 16.
(1905). The Comparative Physiology of the Invertebrate Heart.
Ibid. 13-14.
(1922). A Note on the Action of Drugs on the Invertebrate Heart.
Journ. Gen. Physiol. 4.
Carlson and Meek (1908). On the Mechanism of the Embryonic Heart
Rhythm. Am. Journ. Physiol. 21.
Fredericq (19 1 3). Recherches experimentales sur la physiologic cardiaque
d'Octopus. Arch. Int. Physiol. 14.
Fry (1909). The Influence of the Visceral Nerves on the Heart of Cepha-
lopods. Journ. Physiol. 39.
FucHS (1908). Beitrage zur Physiologic des Kreislaufes bei den Cepha-
lopoden Pflugers Archiv. 60.
Hart (1924). L'action des ions sur les mouvements rhythmiques du sac
muscule cutane du lombric. Arch. Neerland. Physiol. 9.
Hecht (191 8). The Physiology of Ascidia atra. Am. Journ. Physiol. 45.
THE CIRCULATION OF BODY FLUIDS 117
Hoffmann (191 i). Ueber Elektrokardiogram von Evertebraten. Arch
Anat. u. Physiol. 191 1.
Koch (1916). Der Herzschlag von Anodonta. Pflugers Archiv.
156.
LoEB (1903). Ueber die Bedeutung der Blutkorperschen fiir
die Blut-
gerinnung. Virchow's Archiv. 173.
PoLiMANTi (1913). Beitrage zur Physiologic von Maia. Arch. Anat.
u. Physiol. 1 91 3.
Straub (1904). Fortgesetzte Studien am Aplysienherz. Pflugers Archiv
103.
Tait (1918). The Blood of Astacus fluviatilis. Q. E. P. 12.
J.
CHAPTER VII
ENDOCRINE CO-ORDINATION
as a
and the validity of arguing that an effector organ such
melanophore possesses a sympathetic nerve supply from
the
and
by the oesophagus of Aphrodite and Aplysia (Hogben
the heart of Pecten
Hobson) and of Helix (Ten Cate), also
fuscous.
The exact part which light and temperature respectively
play in promoting this sequence of reactions has been made the
subject of investigation by Parker (1906) and Redfield (1918).
From their work it appears that the melanophores of this
Uzard respond to light and darkness, warmth and cold, in the
manner generally characteristic of reptiles, i.e. bright illumina-
tion and low temperature promote darkening of the skin,
while warmth and darkness bring about pallor. Light and
heat interact so that the effect of the latter predominates at
extremes of temperature, and it is thus that, in natural condi-
tions, living as these creaturesdo in a warm climate, pallor
intervenes during that part of the day when the temperature
rises to a maximum.
But in addition to this response to direct illumination,
the horned toad reacts in bodily coloration to the character
of the substratum and to mechanical irritation or disturbance.
Any nocuous stimulus, such as electrical excitation of the roof
of the mouth or the cloaca, evokes pallor in fuscous individuals
ENDOCRINE CO-ORDINATION 125
Background.
Fig. 32. — Frog on right injected six hours previously with'extract of
the pituitary of a foetal ox left, control.
: !
Fig. 33. — Two frogs 19 days after operation on left anterior lobe
:
Further References
ScHAFER. The Endocrine Organs. Longmans, Green.
HoGBEN. The Pigmentary Effector System. Oliver and Boyd.
Swale Vincent. Internal Secretion of the Ductless Glands. Arnold.
CHAPTER VIII
•01 -02
Time since previous stimulus {seconds)
occurring at what was the anode, i.e. at the point to which the
kations now tend to revert. The relation of duration and
intensity of stimulus to the excitation process has been
elucidated chiefly through the work of Lapicque and Keith
Lucas. There exists both a time limit and a limit of intensity
for effective stimulation. If the stimulus is of an intensity
less than a certain amount, cannot excite, however prolonged
it
The
formulae of Nernst account for the experimental data
relatingminimal frequency, duration, and current strength
necessary to set up an excitation within a restricted range.
By taking into consideration the conditions arising when the
membranes at which ions of opposite sign collect are close
together, A. V. Hill deduced a modified expression relating
the duration of an exciting current to its least strength
with greater accuracy than the formulae of Nernst. Hill's
expression is :
_ A
iyiQ^=i —
Thus in an experiment of Lapicque the corresponding
threshold-values for current strength and duration (in o'ooi
sec.) were :
t
. —
140 COMPARATIVE PHYSIOLOGY
To satisfy the above equation the factor /x^* on the left
(1852), before whose time it has been supposed that the nervous
impulse travelled at a rate comparable with the velocity of
—
NERVOUS CONDUCTION AND EXCITATION 141
light ;
and the train of thought which prompted Helmholtz
to experiment is instructive. *'
As long as physiologists thought
itnecessary," he argued, " to refer nerve-actions to the
move-
ments of an imponderable or psychical principle, it
must
have appeared incredible that the velocity of the
movement
could be measured within the short distances of
the body.
At present we know from the researches of du Bois Raymond
upon the electromotive properties of nerve that those activities
by means of which the conduction of an excitation
is
accomplished are in reality actually conditioned by or at least
closely connected with an altered arrangement of their material
particles. Therefore conduction in nerves must belong to
the series of self-propagating reactions of ponderable
bodies.
.
."
. Only six years before Helmholtz' experiment Johannes
Miiller had denied the possibility of ever attaining the means
of measuring the velocity of nervous transmission the history ;
Animal.
142 COMPARATIVE PHYSIOLOGY
In addition to observations on those animals in which it is
Nemertinea i
NERVOUS CONDUCTION AND EXCITATION 143
Alcohol VapOi.
(C:^
Expt. No.
NERVOUS CONDUCTION AND EXCITATION 145
Yic^ 36.
current. The local bio-
electric current leads to
a depolarisation of adjacent parts of the membrane which
in their turn become permeable and the seat of a bio-
electric current. Thus the change is propagated along the
nerve-fibre, the action current in one section becoming the
stimulus for the setting up of a similar condition in the next.
According to this view all stimulation is electrical. It is
of interest therefore to note that incident light produces an
electrical variation (Piper) in the denervated eye both of
Vertebrates and Cephalopods.
The balance of evidence is at present in favour of the view
that in the phenomenon of nervous conduction we have to
deal with a process which is essentially of the same nature
as the excitation process itself. It might therefore be expected
that the capacity of the nerve to conduct a second impulse
NERVOUS CONDUCTION AND EXCITATION 147
Further Reading
Keith Lucas. The Conduction of the Nervous Impulse. Longmans,
Green.
LiLLlE. Protoplasmic Action and Nervous Action. Univ. Chicago Press.
Hill (1921). The Energy Involved in the Electrical Discharge in Muscle
and Nerve. Proc. Roy. Soc, B. 92.
CHAPTER IX
lanql.'oD
Cerebral qanqljon
Preganglionic
neurone
f?ccepfor
Postgangllonrc neurone
Further Reading
Herter. Mechanische Sinnesorgane U. S. W. Leipzig.
LoEB. Forced Movements, Tropisms and Animal Conduct. Lippincott.
that it was not till more than ten years after the issue of the
*'
Origin of Species " that the fertilisation of the egg by a single
sperm was clearly established ;and that the material available
for the study of inheritance by Darwin's contemporaries was
largely derived from popular tradition current among stock-
breeders.
The natural starting-point for a study of the physiology
of reproduction is the fertilisation of the egg. The important
fact that the normal process of fertilisation involves the union
of only one gamete of either sex was first clearly established
by Hertwig and Fol (1875). The recognition of this fact
raises two problems. The entry of the sperm into the egg
normally implies (i) the initiation in the egg of active cell-
division culminating in the formation of a new individual
(2) the transference to the zygote of something in virtue of
which the nev/ individual so formed resembles the male as
to ^8
NaCl. From inspection of the above it is clear that
that this action was due to the acid hydrolysis product. This
suggested that the fatty acids might be successful agents of
membrane formation. By leaving the unfertilised eggs of
Strongylocentrotus in a mixture of 50 c.c. sea water and 2'8
N
c.c. — butyric acid at 15° C. for about two minutes, all the
and the eggs of marine animals are best for this purpose, the
physico-chemical equilibrium being in such cases of a more
mobile character. From the rapid advances made of late
years in the technique of tissue-culture it would not seem
unlikely that the initiation of developmental stages without
contact with sperm will be accomplished in our own time in
mammalian ova.
In the Echinoid egg, which up till now has yielded the most
satisfactory material for experimental manipulation, an im-
portant aspect of the union of the sperm and egg is the
immediate increase in oxygen consumption which occurs
after entry of the sperm. At an early stage in the study of
this problem, Loeb suggested that the immediate effect of
the penetration of the sperm might be to promote a series of
oxidative processes. Warburg's (1908) determinations of the
oxygen-consumption of fertilised and unfertilised eggs of
Arbacia confirmed Loeb's prediction. Warburg found that
a quantity of eggs (about four million) in sea water, equivalent
to 28 mg. by the Kjeldahl estimation, took up
total nitrogen
4-5 c.c. of oxygen during the
first hour after insemination,
later ones the manometer was used for the gas analyses and ;
Egg-water.
THE FERTILISATION OF THE EGG 179
Further Reading
LiLLiE. Problems of Fertilisation. Chicago University Press.
LoEB. Artificial Parthenogenesis and Fertilisation. Ide7n.
INHERITANCE 183
Vestigial, Long-winged.
^/
vv
Vv
VV Vv Vv
Long-winged.
i86 COMPARATIVE PHYSIOLOGY
parent predominates in the heterozygous condition, it is called
the dominant character (long in this case) in contradistinction
to the recessive (vestigial in this example).
An immense variety of characters both in plants and animals
have been found to follow^ the rule of segregation. To mention
but a few, colour of the hair in mam^mals, duration of life in
Drosophila, fecundity and absence of feathers on the neck in
fowls, brachydactyly in man, absence of eyes and wings in
flies. These show what
suffice to diverse types of hereditable
characteristics, anatomical and physiological, depend on
segregating hereditary factors or genes.
However, factorial analysis, as this method of investigation
is sometimes called, is not often as simple as in the case cited.
And those who have criticised the universal applicability of
the gene hypothesis usually do so in the expectation of a text-
book simplicity in eveiy instance. When we cross two strains,
it may, and often does, happen that the difference which
long wings and the same is true if a black fly with vestigial
;
wings is crossed with a fly that is homozygous for the gray body
colour and long- winged condition. But whereas, when the
F.I male from the cross between gray- vestigial and black-
long is mated to the double recessive (black- vestigial) female,
one-half of the off'spring are gray-vestigial and the other half
black-long when the F.i male of the cross between black-
;
but in the F.2, which consists of three reds to one white, all the
females are red- eyed, and all the whites are male. Now when
a pure red-eyed male is crossed with a white-eyed female
the result is quite different ; all the females in the F.i as
before have the dominant red eye ; but the males are white-
eyed. When se^ equal numbers of
the F.i are mated inter
white-eyed and red-eyed females and males are produced.
The inability of the male to transmit red to his offspring of
the same sex is readily explained on the asumption that the
red gene is linked to something which, if present in the zygote
in duplicate, leads to the production of a female, and if present
in the zygote unpaired (diagram) leads to the production of
a male the red-eyed male produces sperm of two kinds, one
;
bearing the " red " gene destined to fertilise an egg which
must become a female, and one which cannot bear the red
gene and which is destined to lead to the production of another
male (Fig, 42). This implies that sex itself is predetermined
1 90 COMPARATIVE PHYSIOLOGY
by genetical factors forwhich one sex is heterozygous, so that
a I I sex ratio is maintained by the normal consequences of a
:
of grossulariata to
ance in Drosophila.
lacticolor, but all the males are of
the grossulariata type. In the reciprocal mating the grossu-
lariata female can only transmit the grossulariata pattern to
her sons ; all the female offspring are of the lacticolor type.
When se^ equal numbers of lacticolor
the F.i are mated inter
and grossulariata males and females are produced (diagram).
Here the female moth is constitutionally simplex with
respect to the sex-Hnked genes. This is the exact reverse of
the state of affairs in sex-linked characters in Drosophila.
Femaleness is associated with the simxplex condition of genes
which in the duplex condition give rise to maleness.
The predetermination of sex by genetic factors does not
.
INHERITANCE 191
races is always the same. In the extreme case all the individuals
of such a cross may be of one sex, but half of these on being
bred back to a parent stock can be shown to have the genetic
constitution of the alternate sex. By making two assumptions
Goldschmidt has brought into line the results of a very large
number of such racial crosses (i): in addition to the genes
for maleness for which the female has the constitution Mw
and the male MM (as in Abraxas where the same type of
sex-linked inheritance occurs) there is something dependent
on the constitution of the tggy transmitted therefore through
the female parent only, that mxodifies the degree of maleness,
and is denoted by the symbol F. Thus a female forms MmF
gametes MF and rriF while the male MMF forms gametes M
only ; (2) it|is^/urther assumed that in different local races the
efficiency of M and F respectively to influence differentiation
in the direction of maleness or femaleness are quantitatively
192 COMPARATIVE PHYSIOLOGY
different. By denoting M
and F in terms of an arbitrary
system of numerical symbols Goldschmidt has elaborated
a system by which the appearance and degree of intersexuality
of his crosses between local races can be faithfully predicted.
Race A T^ac€ B.
Female
^walc
F^ femaUs
CM>F)
INHERITANCE 195
on the one hand and of the dogfish and angel-fish on the other,
differs in this significant respect. They have been laved for
years, or one might properly say for generations before the
experiment, with solutions of different hydrogen ion concentra-
tion."
It is here implied that the action of a stimulus upon the
INHERITANCE 199
Further Reading
GoLDSCHMiDT. Mechanism and Physiology of Sex Deteniiination.
Methuen.
Morgan. The Physical Basis of Heredity. Lippincott.
Crew. Introduction to the Science of Animal Breeding. Oliver and
Boyd.
East and Jones. Inbreeding and Outbreeding. Lippincott.
Morgan, Bridges and Sturtevant (1925). The Genetics of Drosophila.
Bibliographica Genetica (the Hague), vol. 2,
CHAPTER XII
15
— —
THE PHYSIOLOGY OF DEVELOPMENT 207
tions of the germ layer theor^^ For this reason the independent
2o8 COMPARATIVE PHYSIOLOGY
experiments of Shearer have been cited in some detail. It
(2) resorption of the dorsal fin and shedding of the larval skin ;
(3) closure of the gill clefts. The Urodele larva has fully-
THE PHYSIOLOGY OF DEVELOPMENT 209
developed limbs and the tail persists into the adult stage.
The nature of the physiological change which initiates this
series of events is the same in either case.
The first experiment which threw any light on this was
the discovery of Babak (191 1) that the axolotl larva of the
Mexican salamander {Amhlystoma tigrinu7n ) which is nor-
mally neotenous, can be induced to undergo transformation
into the adult form by thyroid administration. Gudernatsch
(19 1 2- 14) showed that this was true of frog tadpoles. If
tadpoles are fed on diets of various tissues, ovary, liver, thymus,
brain, pancreas, spleen, pituitary, and thyroid —those fed on
thyroid gland develop limbs and lose their tails long before
the others. Thus in Rana catesbana (Swingle), a species
which normally requires three seasons to attain to the stage
at which metamorphosis occurs in nature, the six- weeks-old
tadpole will transform into a pigmy frog if fed on ox thyroid.
These observations received abundant confirmation both as
regards urodele larvae and anuran tadpoles (Morse, Barthelemez,
Jensen, Huxley and Hogben, Uhlenhuth).
Bennet Allen (19 16- 18) succeeded in overcoming the
manipulative difficulties of extirpating the thyroid gland in
tadpoles of the toad. The
thyroidectomised tadpoles behave
in a perfectly normal manner until the limb-buds develop,
when transformation should occur. Instead of undergoing
metamorphosis at this stage they remain permanently in the
larval state, attaining as age advances dimensions far exceeding
those of a normal tadpole. They can, however, be induced, as
Swingle (19 18) showed, to develop into normal frogs if fed
on thyroid tissue. Later E. R. and M. M. Hoskins confirmed
the work of Allen by similar experiments on frogs and others
on urodele larvae.
Thus both inAnura andUrodeles it is certain (i) that the
removal of the thryoid normally prevents metamorphosis ;
Further Reading
Child, C. M. (1915). Individuality in Organisms. Chicago.
(1924). Physiological Foundations of Behaviour. New York.
DCrken, B. (1919). Einfiihrung in die Experimentalzoologie. Berlin.
GoLDSCHMiDT, R. (1923). The Mechanism and Physiology of Sex Deter-
mination. London.
Jenkinson, J. W. (1909). Experimental Embryology. Oxford.
Wilson, E. B. (1925). The Cell in Development and Heredity (3rd ed.).
New York.
Recent Papers
Champy (1919). Arch. Morphol. Exp. et Gen., 1919.
Helff, O. M. (1924). The Oxygen Consumption of Thyroid and Diiodo-
tyrosine-fed Tadpoles. Proc. Soc. Exp. Biol. Med., 21, 34.
Huxley, J. S. and de Beer, G. R. (1923). Studies in Dedifferentiation,
IV. Quart. Journ. Micr. Sci., 67, 473.
Huxley, J. S. (1925). Studies in Amphibian Metamorphosis, II. Proc.
Roy. Soc, (B) 98, 113.
INDEX
Abraxas, 190 Baumberger, 87
Absorption spectra, 70, 73, 76, 78 Bayliss, 33, 119
Acids, secretion of, 62 Bernstein, 13, 58, 63, 145
Adler, 130 Bert, 64
Adrenaline, 39, 122 Bethe, 32, 154
Adrian, 14, 136, 143 Biedermann and Moritz, 97
Agglutination, 179 Bioluminescence, 52 et seq.
Aggregation, 179 Blood pressure, 103, 107, 11
Allen, 130, 209 et seq. Bridges, 195
All or nothing law, 143 Briicke, 99, 106
Alsberg and Clark, 75, 105 Bodansky and Rose, 92, 96
Amino acids, 92, 93 Bodo, 66
Amphibian metamorphoses, 208 et Botazzi, 82, 104
seq. Bounhiol, 66
Amphioxus, 90 Robert Boyle, 53
Amoeboid movement, 30 Buchner, 55
Amylolytic enzymes, 96 Buddenbrock and Rohr, 67 et seq.
Anaphylaxis, 59
Anodon, 15, 114
Anson and Mirsky, 79 Calcium ions, 16-22, 27, 35, 163
Antidromic action, 103 Calliphora, 93
Aphrodite, 99, 122 Cancer, 77
Aplysia, 81, 99, 113, 122, 157 Capillaries, 102
Arbacia, 71 Carbon dioxide, tension of, 68, 80
Arenicola, 73, 162 Carbon monoxide, 74
Ascidia, 114 Carcinus, 16, 154
Associative behaviour, 163 Carlson, 107 et seq., iii
Astacus, 16, 77, 97 Carter, 174
Atlantic minnow, 20, 38, 201 Catch muscle, 15
Atzvell, 132 Cellulose, 97
Aurelia, 153 Cephalopods, eye of, 146
Autonomic ganglia, 157 nervous system, 155
Axial gradients, 203 Cerianthus, 152
Axolotl, 209 et seq. Ceriodaphnia, no
Chaetopterus, 52
Chambers, 35, 152
Babak, 69 Chemotaxis of sperm, 178-9
Bacot and Harden, 88 Child, 201 et seq.
Baglioni, 52 Chironomus, 65
Banting and Best, 98 Chlamydomonad, 89
Barcroft, 28, 49, 50, 73, loi, 102, Chlorhaemidse, 77
175 Chlorocruorin, 77 et seq.
Barium, 39 Chromatophores of Crustacea, 39
Bateson, 183 Chromosomes, 193 et seq.
215
2l6 INDEX
Chronaxie, 137 Ergotoxine, 39
Ciliary feeding, 89-91 Esculin, 56
motion, 23 et scq., 151 Lovatt Evans, 16
Ciona, 180 Excitation in nerve, 135 et scq.
Enzymes, 54 et seq. i
Hibernation, 95
1
INDEX 217
2l8 INDEX
of 28-9 Reversal of rhythm in tunicate
Oxygen consumption cilia,
heart, 115
of eggs, 17s
embryos, 206 Rhumhler, 33
muscle, 8-1 Ritchie, 8
Oxy haemoglobin, 70 Roaf, 123
Robber fly, 160
Robertson, Brailsford, no
Palaemon, 42 Rogers, 132
Palinurus, 81, 107 Romanes, 153
Pancreas, 97, 119 Rouget, cells of, 102
Pantin, 32 et seq.
Parker, 124, 152, 161
Parnas, 15 Sabelliformia, 77
Parsons, 81 Sanford, 94
Pasteur, 55 Schafer, 121
Pathenogenesis, i']i et seq, Schulz, 61
Pazvlozv, 164 et seq. Scy Ilium, 17
Pecten, 15 Secretion, 119
Pedicellariae, 59
Self-sterility, 180
Pepsin, 92 Sepia, 53
Peters, 12, 71
Sergester, 53
Pfluger's Laio, 137
Sex determination, 189-93
differentiation, 211
Phallusia, 81
Pholas, 53 Shearer, 175, 206
Phototropism, 159 e^ ^^3- Sherrington, 147, I49> 158
Phrynosoma, 124 Simulium, 66
Phyllirhoe, 53 Smith, P. E., 120, 130
Physalia, 92 Snail, 17, 77, 97
Physoclisti, 62 Spftth, 38 ei seq.
Pigmentary effect on system, 37 e/ Spallanzani, 53
seq. Sperographis, 77
Pineal, 133 Sponges, 151
Piper, 146 Stannius experiment, 112
Pituitary gland, 103, 127, 211 Starling, 49
Pollack, 63 Statoliths, 156
Porthesia, 159, 163 Stedman, 76
Pouchet, 120 Stick insect, 67
Protacanthus, 160 Stigmata, 67
Purpura, 123 Stockard, 201
Putter's hypothesis, 89 Strauh, 113
Pyrosoma, 53 Stretching, effect of, on muscle, 113
Pyrophorus, 53 Strongylocentrotus, 172
Strong acids and bases, 25-6
Suprarenals, 127
Quagliariello, 75 > ^3 Susceptibility method, 204
Syllid, 99
Synoptic nervous system, 154
Raia, 17 Swingle, 131, 209
Recessive, 186
Redfield, 124
Refractory period, 135 Takamine, 121
Renal secretion, 47 et seq. Teleosts, 97,
tubules, 48 blood of, 104
Rennet, 54 Temperature, 28-9, no
Rennie, 91 Ten Gate, 99, 114, 122
Respiratory movements, 67, 69 Tentaculocysts, 153
quotient, 30, 49, 95 Tetanus, 3
1 1
INDEX 219
Thyroid gland, 209 et seq. Van 't Hoff solution, 25
Tonus, 15 V. der Heyde, 5
Torpedo, 58 Venoms, 60
Tracheal respiration, 66 et seq, Vitamins, 88
Trivalentious, 18 Vies, 74
Tropisms, 158 Voit, 85
Tunicate heart, 114 V, Frisch, 120
Tyramine, 60, 122
Yeasts, 87-9
Vanadium, no I
I
Yonge, 92, 97
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